Biodiversity of Inland Waters

**152**

*Inland Waters - Dynamics and Ecology*

[1] Sun F, Sun W, Chen J, Gong P. Comparison and improvement of methods for identifying water bodies in remotely sensed imagery. International

[9] Jiang H, Feng M, Zhu Y. An

Sensing. 2014;**6**(6):5067-5089

extraction for remote sensing images from urban area based on morphological transformations. IEEE Transaction on Geosciences and Remote

Sensing. 2003;**41**(9):1940-1949

[12] Wu B, Fang C, Yu L, Huang X, Zhang Q. A fully automatic method to extract rare earth mining area from Landsat image. Photogrammetric Engineering and Remote Sensing.

[13] Huang X, Zhang L, Wang L. Evaluation of morphological texture features for mangrove forest mapping and species discrimination using multispectral IKONOS imagery. IEEE Geoscience and Remote Sensing Letters.

[14] Ji L, Zhang L, Wylie B. Analysis of dynamic thresholds for the normalized difference water index. Photogrammetric Engineering and Remote Sensing. 2009;**75**(11):1307-1317

[15] Otsu N. A threshold selection method from gray-level histograms. IEEE Transactions on Systems, Man and

Cybernetics. 1979;**9**(1):62-66

[16] Zha Y, Gao Y, Ni S. Use of

normalized difference built-up index in automatically mapping urban areas from TM imagery. International Journal of Remote Sensing. 2003;**24**(3):583-594

[11] Huang X, Zhang L. A multidirectional and multi-scale morphological index for automatic building extraction from multispectral GeoEye-1 imagery. Photogrammetric Engineering & Remote Sensing.

2011;**77**(7):721-732

2016;**82**(9):55-64

2009;**6**(3):393-397

[10] Beneditsson JA, Pesaresi M, Arnason K. Classification and feature

automated method for extracting rivers and lakes from Landsat imagery. Remote

[2] Frazier PS, Page KJ. Water body detection and delineation with Landsat TM data. Photogrammetric Engineering and Remote Sensing.

[3] Rokni K, Ahmad A, Solaimani K, Hazini S. A new approach for surface water change detection: Integration of pixel level image fusion and image classification techniques. International Journal of Applied Earth Observation and Geoinformation. 2015;**34**:226-234

[4] McFeeters S. The use of the normalized difference water index (NDWI) in the delineation of open water features. International Journal of Remote Sensing. 1996;**17**:1425-1432

[5] Xu H. Modification of normalised difference water index (NDWI) to enhance open water features in remotely sensed imagery. International Journal of Remote Sensing. 2006;**27**:3025-3033

[6] Carleer AP, Wolff E. Urban land cover multi-level region-based classification of VHR data by selecting relevant features. International Journal of Remote Sensing.

2006;**27**(6):1035-1051

2010;**39**(6):611-617

[7] Yang S, Xue S, Liu T. Method for automatically extracting fine water body by using TM image. Journal of Surveying and Mapping (in Chinese with English Abstract).

[8] Li Y, Ding J, Yan R. Study on extraction method of small water bodies in mountainous area based on GF-1 remote sensing images. Resource Science (in Chinese with English Abstract). 2015;**37**(2):408-416

Journal of Remote Sensing.

2012;**33**:6854-6875

**References**

2000;**66**:1461-1468

**Chapter 10**

**Abstract**

**1. Introduction**

arbitrarily small size category.

**155**

*Kenneth W. Cummins*

Stream Invertebrate Zoology

an avalanche of running water invertebrate research.

For over a century, there has been strong interest in freshwater streams and rivers. Since the inception of studies on running waters, invertebrates have been a central theme. Early descriptive work in Scandinavia and New Zealand was followed by work in Europe, England, and then North America and Australia. Presently, there is a very significant interest worldwide including Asia, Central and South America, and Africa in freshwater invertebrates. Throughout, insects have dominated the focus on invertebrates. Although the major marine invertebrate groups are present in freshwaters, there are essentially no marine insects. A clear picture of the habitat and food requirements of running water invertebrates shows that they serve as important indicators of water quality and fisheries. Major paradigms, such as the River Continuum and functional feeding groups, have provided frameworks for studies of running water (lotic) invertebrates. Once stream and river research achieved an international status by separation from lake domination of the limnology discipline, there has been

**Keywords:** stream and river ecology, lotic invertebrates, functional feeding

by the time insects evolved, all the marine ecological niches were filled.

Macroinvertebrates conventionally have been defined as those individuals greater than 1 mm in size. However, many present-day studies include all invertebrates retained on a 0.25 -mm mesh screen as macroinvertebrates (1). The far less studied and much smaller microinvertebrates include taxa also found in the marine ecosystem such as protozoans, rotifers, and annelids. But, some very small insect taxa (Diptera, *Chironomidae*) and the first instar of most aquatic insects are also in this

Examples of earlier investigations of running water invertebrates can be found in Shelford [4] and Shelford and Edy [5] (North America), Moon [6, 7] (Great Britain), Wessenberg-Lund [8] (Denmark), and Allen [9] (New Zealand). The North American references [4, 5] contain early descriptive work on the components of lotic invertebrate populations and their habitats. The British publication [6]

For over a century, there has been significant interest in stream and river (lotic) ecology. A major foundation fueling this interest has been the aquatic invertebrates. From the beginning, focus has been on certain marine-derived groups and on insects. Lotic macroinvertebrate communities are usually dominated by insects [1], but some marine taxa, such as annelids, mollusks, and crustaceans, are often abundant as well [2, 3]. There are essentially no marine insects, the argument being that

groups (FFG), River Continuum, FFG ecosystem surrogate ratios

#### **Chapter 10**

### Stream Invertebrate Zoology

*Kenneth W. Cummins*

#### **Abstract**

For over a century, there has been strong interest in freshwater streams and rivers. Since the inception of studies on running waters, invertebrates have been a central theme. Early descriptive work in Scandinavia and New Zealand was followed by work in Europe, England, and then North America and Australia. Presently, there is a very significant interest worldwide including Asia, Central and South America, and Africa in freshwater invertebrates. Throughout, insects have dominated the focus on invertebrates. Although the major marine invertebrate groups are present in freshwaters, there are essentially no marine insects. A clear picture of the habitat and food requirements of running water invertebrates shows that they serve as important indicators of water quality and fisheries. Major paradigms, such as the River Continuum and functional feeding groups, have provided frameworks for studies of running water (lotic) invertebrates. Once stream and river research achieved an international status by separation from lake domination of the limnology discipline, there has been an avalanche of running water invertebrate research.

**Keywords:** stream and river ecology, lotic invertebrates, functional feeding groups (FFG), River Continuum, FFG ecosystem surrogate ratios

#### **1. Introduction**

For over a century, there has been significant interest in stream and river (lotic) ecology. A major foundation fueling this interest has been the aquatic invertebrates. From the beginning, focus has been on certain marine-derived groups and on insects. Lotic macroinvertebrate communities are usually dominated by insects [1], but some marine taxa, such as annelids, mollusks, and crustaceans, are often abundant as well [2, 3]. There are essentially no marine insects, the argument being that by the time insects evolved, all the marine ecological niches were filled. Macroinvertebrates conventionally have been defined as those individuals greater than 1 mm in size. However, many present-day studies include all invertebrates retained on a 0.25 -mm mesh screen as macroinvertebrates (1). The far less studied and much smaller microinvertebrates include taxa also found in the marine ecosystem such as protozoans, rotifers, and annelids. But, some very small insect taxa (Diptera, *Chironomidae*) and the first instar of most aquatic insects are also in this arbitrarily small size category.

Examples of earlier investigations of running water invertebrates can be found in Shelford [4] and Shelford and Edy [5] (North America), Moon [6, 7] (Great Britain), Wessenberg-Lund [8] (Denmark), and Allen [9] (New Zealand). The North American references [4, 5] contain early descriptive work on the components of lotic invertebrate populations and their habitats. The British publication [6]

established the fundamental classification of flowing water habitats as either erosional (riffles) or depositional (pools). This basic view endures to the present time and is similar to the lake (lentic) designations of littoral or profundal [10]. The basic morphological and behavioral adaptations of lotic invertebrates, to either erosional or depositional habitats, are discussed below. The Scandinavian volume by Wessenberg-Lund contains a treasure trove of biological and ecological information on freshwater insects including habitat descriptions [8]. Someone could contribute significantly to the study of lotic invertebrates by translating this book from Danish German into English. The famous New Zealand publication by Allen [9] used unique illustrations of the fauna to represent relative densities of the stream macroinvertebrates. A wide-ranging geographical scope of lotic invertebrate study was, and is, an important component of the broad development of the field of stream and river ecology. The discussion on running water invertebrates that follows is greatly informed by recent advances in taxonomy, biology, and ecology [2, 3], especially of insects. Examples are the new (5th) edition of the aquatic insects of North America (Berg et al. [1]), DNA barcoding to validate taxonomic affinities [11, 12], stable isotope (carbon 13, nitrogen 15) analysis of food webs [13, 14], and functional feeding group (FFG) characterization of trophic relationships [15]. It is now possible to identify most genera of North American aquatic insects using morphological characters [1]. It should be noted that many of the North American families and genera occur on other continents.

**Taxa Primary energy source;**

*DOI: http://dx.doi.org/10.5772/intechopen.88283*

**Oligochaeta** (segmented worms)

*Stream Invertebrate Zoology*

**Gastropoda** (snails)

**Crustacea** Amphipoda, *Gammarus* (scuds) [22], Isopoda, *Assellus* (*sow bugs*)

**Ephemeroptera** (mayflies) Heptageniidae, Ephemerellidae, *Drunella*

Baetidae, *Baetis* Leptophlebiidae, *Paraleptophlebia*

Ephemeridae *Ephemera*

**Trichoptera** (caddisflies) Limnephilidae<sup>1</sup>

*Hydatophylax*, *Pycnopsyche*

Hydropsychidae, Philopotamidae

**157**

,

**and habitat**

Autochthonous FPOM produced by invertebrate feeding, mechanical breakage of CPOM, and from the stream bank, pools, backwaters, and margins

Autochthonous attached nonfilamentous algae; in riffles

Allochthonous CPOM riparian plant litter; accumulations against obstructions, in backwaters, pools, and stream margins

Autochthonous attached nonfilamentous algae; in riffles

Autochthonous FPOM produced by invertebrate feeding, mechanical breakage of CPOM, and from the stream bank, pools, backwaters, and margins

Autochthonous produced by invertebrate feeding, mechanical breakage of CPOM, and from the stream bank; gravel riffles

Allochthonous riparian plant litter; accumulations against obstructions in the current and in backwaters or pools

CPOM, FPOM, and small invertebrates in the appropriate size that can be retained in filtering nets; retreats fastened to coarse substrate in riffles

algae

invertebrate prey; riffles or plant litter accumulations

Hydroptilidae Autochthonous filamentous

Rhyacophilidae Autochthonous in-stream

**Food resource category Habit (mode of**

FPOM (organic particles, coated and colonized by bacteria, including invertebrate feces); deposited on or in the bottom sediments

Periphyton: attached nonfilamentous algae and associated FPOM and microinvertebrates

Conditioned (colonized by microbes, especially aquatic hyphomycete fungi) CPOM plant litter

Periphyton: attached nonfilamentous algae and associated FPOM and microinvertebrates

FPOM (organic particles, coated and colonized by bacteria, including invertebrate feces); deposited on or in the bottom sediments

FPOM (organic particles), including invertebrate feces, coated and colonized by bacteria in the size range that can be pumped through burrows

Conditioned (colonized by microbes, especially aquatic hyphomycete fungi) CPOM plant litter

FPOM (organic particles, including invertebrate feces, coated and colonized by bacteria and small invertebrates in the appropriate size that is caught by capture nets)

Individual filamentous algal cell contents

> Invertebrate prey of appropriate size

**attachment, concealment, movement)**

Burrowers, mostly in fine sediments

Clingers, on coarse sediments in riffles

Burrowers, in plant litter accumulations

Clingers, on coarse sediments in riffles

Swimmers, in pools, backwaters, and margins, occasionally moving through riffles

Burrowers, in gravel riffles where they pump water through burrows

Burrowers, in plant litter accumulations

Clingers, on coarse sediments in riffles

Climbers, in filamentous algal colonies

Clingers on coarse sediments or sprawlers in plant litter accumulations **FGG**

**GC**

SC

DSH

SC

GC

FC

DSH1 (SC)

FC

PC

P

#### **2. Taxonomic invertebrate groups of lotic invertebrates**

#### **2.1 Macroinvertebrates**

The typical macroinvertebrate groups and their characteristics in headwater streams are summarized in **Table 1**. The conterminous continent-wide US study of selected river basins, the River Continuum Project [16], developed a paradigm linking position along a river basin channel network that was described as stream order [17]. Energy sources for the component communities of macroinvertebrates were predicted along the continuum. Headwater streams (orders 1–3) receive their energy supply from streamside (riparian) terrestrial vegetation (plant litter) along the stream channel. This is termed an allochthonous energy source.

Wider mid-sized stream/rivers are less shaded by riparian trees, allowing much more light to reach the channel. This additional light input drives in-stream primary production by algae and aquatic vascular plants. This is termed an autochthonous energy source. The macroinvertebrate communities of the mid-sized stream/river ecosystems are populated with taxa that utilize the greatly increased beds of rooted aquatic vascular plants, especially herbivore shredders. Also, these order 4–6 running waters have fauna which contains headwater taxa derived from terrestrial ancestors (insects) together with taxa of marine ancestral origin (e.g. mollusks) (**Table 2**) [16].

The dominant energy source for larger rivers (orders 7–10 or greater) is input from the upstream channel network (orders 1–7) plus periodic return flow from the floodplain [16, 18–21]. These larger rivers are usually turbid, and, although adequate light reaches the surface of the water, penetration to the bottom is poor and primary production is restricted. The dominant food resource for the invertebrates is fine particulate organic matter (FPOM). FPOM consists of particles >1 mm in size, organic and mineral particles surface-colonized by bacteria, and algae and microinvertebrates in suspension. The primary mode of feeding for the

established the fundamental classification of flowing water habitats as either erosional (riffles) or depositional (pools). This basic view endures to the present time and is similar to the lake (lentic) designations of littoral or profundal [10]. The basic morphological and behavioral adaptations of lotic invertebrates, to either erosional

Wessenberg-Lund contains a treasure trove of biological and ecological information on freshwater insects including habitat descriptions [8]. Someone could contribute significantly to the study of lotic invertebrates by translating this book from Danish German into English. The famous New Zealand publication by Allen [9] used unique illustrations of the fauna to represent relative densities of the stream macroinvertebrates. A wide-ranging geographical scope of lotic invertebrate study was, and is, an important component of the broad development of the field of stream and river ecology. The discussion on running water invertebrates that follows is greatly informed by recent advances in taxonomy, biology, and ecology [2, 3], especially of insects. Examples are the new (5th) edition of the aquatic insects of North America (Berg et al. [1]), DNA barcoding to validate taxonomic affinities [11, 12], stable isotope (carbon 13, nitrogen 15) analysis of food webs [13, 14], and functional feeding group (FFG) characterization of trophic relationships [15]. It is now possible to identify most genera of North American aquatic insects using morphological characters [1]. It should be noted that many of the North American

or depositional habitats, are discussed below. The Scandinavian volume by

families and genera occur on other continents.

*Inland Waters - Dynamics and Ecology*

**2.1 Macroinvertebrates**

(**Table 2**) [16].

**156**

**2. Taxonomic invertebrate groups of lotic invertebrates**

the stream channel. This is termed an allochthonous energy source.

The typical macroinvertebrate groups and their characteristics in headwater streams are summarized in **Table 1**. The conterminous continent-wide US study of selected river basins, the River Continuum Project [16], developed a paradigm linking position along a river basin channel network that was described as stream order [17]. Energy sources for the component communities of macroinvertebrates were predicted along the continuum. Headwater streams (orders 1–3) receive their energy supply from streamside (riparian) terrestrial vegetation (plant litter) along

Wider mid-sized stream/rivers are less shaded by riparian trees, allowing much more light to reach the channel. This additional light input drives in-stream primary production by algae and aquatic vascular plants. This is termed an autochthonous energy source. The macroinvertebrate communities of the mid-sized stream/river ecosystems are populated with taxa that utilize the greatly increased beds of rooted aquatic vascular plants, especially herbivore shredders. Also, these order 4–6 running waters have fauna which contains headwater taxa derived from terrestrial ancestors (insects) together with taxa of marine ancestral origin (e.g. mollusks)

The dominant energy source for larger rivers (orders 7–10 or greater) is input from the upstream channel network (orders 1–7) plus periodic return flow from the floodplain [16, 18–21]. These larger rivers are usually turbid, and, although adequate light reaches the surface of the water, penetration to the bottom is poor and primary production is restricted. The dominant food resource for the invertebrates is fine particulate organic matter (FPOM). FPOM consists of particles >1 mm in size, organic and mineral particles surface-colonized by bacteria, and algae and

microinvertebrates in suspension. The primary mode of feeding for the



**Taxa Primary energy source;**

*DOI: http://dx.doi.org/10.5772/intechopen.88283*

Oligochaeta (segmented worms)

*Stream Invertebrate Zoology*

Bivalvia (bivalve clams) *Sphaerium, Pisidium*

Crustacea Decapoda (crayfish)

Crustacea Amphipoda, *Hyalella*

Ephemeroptera Ephemeridae

Ephemeroptera Baetidae

Hemiptera Corixidae

Ephemeroptera Baetidae

Trichoptera Hydropsychidae

*[1, 3, 19, 22, 26, 28, 29].*

**Table 2.**

**159**

**habitat**

Autochthonous FPOM produced by invertebrate feeding, mechanical breakage of CPOM, and from the stream bank; gravel riffles

Autochthonous FPOM (transport from upstream and from river banks, invertebrate feces, and mechanical breakage of CPOM); bottom sediments

CPOM detritus; rooted aquatic plant beds, pools, backwaters, side channels, and river margins where CPOM detritus accumulates

Autochthonous rooted aquatic plant beds; stems of rooted plants

Autochthonous FPOM produced by invertebrate feeding, mechanical breakage of CPOM from bank and riffles

Autochthonous FPOM, settled in depositional areas, especially rooted vascular plant beds

Autochthonous rooted aquatic plant beds; stems of rooted plants

Autochthonous FPOM, settled in depositional areas, especially rooted vascular plant beds

Autochthonous cell contents of filamentous algae

*Typical North American macroinvertebrates of mid-sized rivers (orders 4–6).*

**Food resource category Habit (mode of**

FPOM particle in the bottom sediments

FPOM organic particles of appropriate size to be filtered through incurrent siphon of material in transport at water sediment interface

Fragmenting and decomposing rooted vascular plant tissue (and some live vascular plant tissue)

Periphyton (algae and associated detritus and microarthropods on rooted plant stems)

FPOM in the size range that can be pumped through burrows

FPOM (organic particles, coated and colonized by bacteria, including invertebrate feces)

> Periphyton algae on rooted plant stems

FPOM (organic particles, coated and colonized by bacteria, including invertebrate feces)

Individual cell contents of filamentous algae

*Allochtonous is energy source from outside the stream channel (riparian zone); autochthonous is energy derived within the stream; CPOM is coarse particulate organic matter >1 mm size; FPOM is fine particulate organic matter <1 mm size; conditioned CPOM is riparian plant litter (e.g. leaves and needles); conditioning involves colonization by microbes, especially aquatic hyphomycete fungi; FFG is functional feeding group: SC = scrapers; GC = gathering collectors; FC = filtering collectors; DSH = detrital shredders, HSH = herbivore shredders; PC = algal cell piercers; P = predators*

**attachment, concealment, or movement)**

Burrowers in the sediments

Burrowers (with incurrent siphon above sediment surface to allow for filtering of FPOM)

Sprawlers (in accumulations of CPOM surface and rooted plant beds)

Climbers (on rooted aquatic vascular plant stems)

Burrowers in river bed gravel, sand, mud sediments with sufficient flow to provide FPOM to be pumped through burrow tube

Swimmers (among rooted aquatic plant beds and backwaters)

Climbers (on rooted aquatic vascular plant stems)

Swimmers (among rooted aquatic plant beds and backwaters)

**FFG**

GC

FC

DSH HSH

SC

FC

GC

SC

GC

FC

*Allochtonous energy has source from outside the stream channel (riparian zone); autochthonous energy source within the stream; CPOM is coarse particulate organic matter >1 mm size [23]; FPOM is fine particulate organic matter <1 mm size [24]. Conditioned CPOM is riparian plant litter (e.g. leaves and needles); conditioning involves colonization by microbes, especially aquatic hyphomycete fungi [25]; FFG is functional feeding group: SC = scrapers, DSH = detrital shredders; HSH herbivore Shredders, GC = gathering collectors; FC = filtering collectors, PC = algal cell piercers, P = predators [1, 16, 19, 22, 26–28]. <sup>1</sup>*

*Many genera have organic cases in the first four instars and are DSH but have mineral cases in the 5th instar and are scrapers.*

#### **Table 1.**

*Typical North American macroinvertebrates of headwater streams (orders 1–3) [22, 27, 29].*

**Taxa Primary energy source;**

*Inland Waters - Dynamics and Ecology*

**Coleoptera** (beetles) Psephenidae (water pennies) Elmidae (adults)

Dytiscidae (larvae and adults) Gyrinidae (larvae and adults) Hydrophilidae (larvae)

Hydrophilidae (adults)

**Diptera** (true flies)

Tipulidae,*Tipula* Orthocladiinae, *Brillia*

Chironomidae (midges) Chironomini Orthocladiinae genera

Simuliidae (blackflies)

Chironomidae Tanypodinae Ceratopogonidae (biting midges, no-see-ums)

*22, 26–28]. <sup>1</sup>*

*scrapers.*

**Table 1.**

**158**

**and habitat**

Autochthonous attached nonfilamentous algae; in riffles

Autochthonous macroinvertebrate prey in pools and backwaters

Autochthonous FPOM detritus settled in pools and backwaters

Allochthonous plant litter accumulations against obstructions, in backwaters, pools, and at stream margins

Autochthonous FPOM deposited in sediments in pools, backwaters, and slow riffles

transport in habitats with moderate flow

Autochthonous clingers; coarse sediments, or wood in riffles

Autochthonous small invertebrate prey

Tanytarsini Autochthonous FPOM in

**Food resource category Habit (mode of**

Periphyton: attached nonfilamentous algae and associated FPOM and microinvertebrates

> Invertebrate prey of appropriate size

FPOM organic particles plus microbes

Conditioned (colonized by microbes, especially aquatic hyphomycete fungi) CPOM plant litter

FPOM (organic particles, including invertebrate feces, coated and colonized by bacteria); deposited on or in the bottom sediments

FPOM (organic particles), including invertebrate feces, coated and colonized by bacteria of appropriate size to be captured

FPOM (organic particles), including invertebrate feces, coated and colonized by bacteria in the size range that can be captured by the filtering head fans; suspended in the passing water column

Small prey (e.g. midges, blackflies)

*Allochtonous energy has source from outside the stream channel (riparian zone); autochthonous energy source within the stream; CPOM is coarse particulate organic matter >1 mm size [23]; FPOM is fine particulate organic matter <1 mm size [24]. Conditioned CPOM is riparian plant litter (e.g. leaves and needles); conditioning involves colonization by microbes, especially aquatic hyphomycete fungi [25]; FFG is functional feeding group: SC = scrapers, DSH = detrital shredders; HSH herbivore Shredders, GC = gathering collectors; FC = filtering collectors, PC = algal cell piercers, P = predators [1, 16, 19,*

*Many genera have organic cases in the first four instars and are DSH but have mineral cases in the 5th instar and are*

*Typical North American macroinvertebrates of headwater streams (orders 1–3) [22, 27, 29].*

**attachment, concealment, movement)**

Clingers, on coarse sediments in riffles

Sprawlers (larvae), swimmers (adults)

Swimmers, in pools and backwaters

Burrowers, in plant litter accumulations

Burrowers, in fine sediments

Clingers, on substrates in moderate current

Clingers, on coarse sediments in riffles

Clingers, on coarse sediments in riffles or in plant litter accumulations

**FGG**

SC

P

GC

DSH

GC

FC

FC

P


*Allochtonous is energy source from outside the stream channel (riparian zone); autochthonous is energy derived within the stream; CPOM is coarse particulate organic matter >1 mm size; FPOM is fine particulate organic matter <1 mm size; conditioned CPOM is riparian plant litter (e.g. leaves and needles); conditioning involves colonization by microbes, especially aquatic hyphomycete fungi; FFG is functional feeding group: SC = scrapers; GC = gathering collectors; FC = filtering collectors; DSH = detrital shredders, HSH = herbivore shredders; PC = algal cell piercers; P = predators [1, 3, 19, 22, 26, 28, 29].*

**Table 2.**

*Typical North American macroinvertebrates of mid-sized rivers (orders 4–6).*


microinvertebrates (zooplankton) and macroinvertebrates (e.g. clams) of the rivers

*Allochtonous is energy source7–10i derived from upstream channel network (orders 1–6); FFG i = functional feeding group: SC = scrapers; GC = gathering collectors; FC = filtering collectors; DSH = detrital shredders, HSH = herbivore*

The very small lotic microinvertebrate taxa are shared with marine environments, such as Protozoa, Rotifera, Nematoda, and micro-Crustacea (Cladocera and Copepoda) [32]. The unofficial definition of microinvertebrates is individuals smaller than 1 mm. Defined this way, early stages of macroinvertebrates need to be included. The difference is that "true" microinvertebrates do not grow beyond the 1 mm size-defined category. In the vast majority of studies in running waters (almost exclusively in rivers), the term microinvertebrates can be replaced by zooplankton. Zooplankton are small invertebrates that live suspended in the water column and have limited ability to control their location [2]. Studies of running water macroinvertebrates vastly exceed those of zooplankton. The River Continuum project [16] recognized zooplankton as a dominant group in rivers of orders 7–10 together with benthic macroinvertebrate such as oligochaetes, bivalve gastropods, and some micro-crustaceans [1]. Studies of river zooplankton have focused on their role in food chains of fish and organic matter cycling. Although most of the zooplankton inhabit the water column of the river where they filter feed on suspended FPOM [32] the benthic forms filter FPOM from the water column and

Thus, the River Continuum model predicts that the small headwater streams will be dominated by invertebrate taxa that are dependent on an allochthonous energy [16]. The most common macroinvertebrates are Detrital Shredders utilizing coarse particulate organic matter (CPOM) plant litter: (DSH) scuds (Amphipoda), several stonefly (Plecoptera), and caddisfly (Trichoptera) families and crane flies *Tipula* (Diptera,*Tipulidae*) are the macroinvertebrates that feed on the plant litter inputs. Certain mayflies (Ephemeroptera) and midges (Diptera, *Chironomidae*) supported by FPOM generated by FFG DSH taxa. Macroinvertebrates of mid-sized rivers utilize autochthonous food resources, especially aquatic plants. Large river invertebrate food chains depend on autochthonous FPOM in suspension delivered from the

FPOM consisting of organic particles colonized by microbes in suspension

**Food resource category Habit (mode of**

**attachment, concealment, or movement)**

Clingers, on the surface of large woody debris

**FFG**

FC

is filtering (filtering collectors) (**Table 3**) [16].

**Taxa Primary energy source;**

*DOI: http://dx.doi.org/10.5772/intechopen.88283*

*Stream Invertebrate Zoology*

Tanytarsini Autochthonous FPOM in

**Table 3.**

**habitat**

sediments and crevices on large woody debris

transport in the water column from upriver tributaries past large woody debris surfaces along river bank or against point bars

*shredders; PC = algal cell piercers; P = predators [1, 16, 19, 20, 30, 31].*

*Typical North American macroinvertebrates of large rivers (orders 7–10).*

upstream channel network and the floodplain.

**2.2 Microinvertebrates (zooplankton)**

on depositional FPOM [1, 2, 24].

**161**


*Allochtonous is energy source7–10i derived from upstream channel network (orders 1–6); FFG i = functional feeding group: SC = scrapers; GC = gathering collectors; FC = filtering collectors; DSH = detrital shredders, HSH = herbivore shredders; PC = algal cell piercers; P = predators [1, 16, 19, 20, 30, 31].*

#### **Table 3.**

**Taxa Primary energy source;**

*Inland Waters - Dynamics and Ecology*

Oligochaeta (segmented worms)

Bivalvia (large bivalve clams)

**Crustacea** (zooplankton) Cladocera, Copepoda

**Megaloptera** (Dobsonflies) Corydalidae

**Trichoptera** Hydropsychidae

**Coleoptera** (beetles) Dytiscidae, Gyrinidae (larvae and adults), Hydrophilidae (larvae)

Hydrophilidae (adults)

**Chironomidae** (midges) Chironomini

**160**

**Diptera** Simuliidae (blackflies) **habitat**

Autochthonous FPOM transported from upriver tributaries and settled on or trapped in the bottom sediments

Autochthonous FPOM transport from upriver tributaries in the water column past the bottom sediments where the clams reside

Autochthonous phytoplankton, bacteria, rotifers, and protozoans produced in situ and micro- FPOM in the water column

Autochthonous invertebrate prey on large woody debris along river bank or against point bars

Autochthonous FPOM in transport in the water column from upriver tributaries in the water column past the capture nets on large woody debris where the larval retreats are attached

Autochthonous macroinvertebrate prey in pools and backwaters

Autochthonous FPOM detritus plus microbes in backwaters

Autochthonous FPOM in transport in the water column from upriver tributaries past boulders and large cobbles in rapids and large woody debris surfaces

Autochthonous FPOM in from upriver tributaries and deposited in the

**Food resource category Habit (mode of**

FPOM benthic organic particles colonized by microbes

FPOM consisting of organic particles colonized by microbes and phytoplankton and zooplankton in suspension

FPOM consisting of phytoplankton, bacteria, rotifers, and protozoans and microorganic particles

Prey consisting of micro- and macroinvertebrates cohabiting large woody debris (e.g. Diptera Chironomidae)

FPOM consisting of organic particles colonized by microbes and phytoplankton and zooplankton in suspension

Invertebrate prey of appropriate size

FPOM detritus consisting of dead organic matter plus microbes

FPOM consisting of organic particles colonized by microbes in suspension

FPOM consisting of deposited organic particles colonized by microbes

**attachment, concealment, or movement)**

Burrowers, in the sediments

Burrowers, in sediments with siphon above the surface allowing capture and filtration of FPOM in transport

Swimmers, in the water column (limited directed movement, easily carried by any current)

> Clingers, on large woody debris

> Clingers, on large woody debris

Sprawlers (larvae), swimmers (adults), in backwaters

> Swimmers, in backwaters

Clingers, in rapids and on large woody debris

Burrowers, in sediments and crevices in large woody debris

**FFG**

GC

FC

FC

P

GC

P

GC

FC

GC

*Typical North American macroinvertebrates of large rivers (orders 7–10).*

microinvertebrates (zooplankton) and macroinvertebrates (e.g. clams) of the rivers is filtering (filtering collectors) (**Table 3**) [16].

Thus, the River Continuum model predicts that the small headwater streams will be dominated by invertebrate taxa that are dependent on an allochthonous energy [16]. The most common macroinvertebrates are Detrital Shredders utilizing coarse particulate organic matter (CPOM) plant litter: (DSH) scuds (Amphipoda), several stonefly (Plecoptera), and caddisfly (Trichoptera) families and crane flies *Tipula* (Diptera,*Tipulidae*) are the macroinvertebrates that feed on the plant litter inputs. Certain mayflies (Ephemeroptera) and midges (Diptera, *Chironomidae*) supported by FPOM generated by FFG DSH taxa. Macroinvertebrates of mid-sized rivers utilize autochthonous food resources, especially aquatic plants. Large river invertebrate food chains depend on autochthonous FPOM in suspension delivered from the upstream channel network and the floodplain.

#### **2.2 Microinvertebrates (zooplankton)**

The very small lotic microinvertebrate taxa are shared with marine environments, such as Protozoa, Rotifera, Nematoda, and micro-Crustacea (Cladocera and Copepoda) [32]. The unofficial definition of microinvertebrates is individuals smaller than 1 mm. Defined this way, early stages of macroinvertebrates need to be included. The difference is that "true" microinvertebrates do not grow beyond the 1 mm size-defined category. In the vast majority of studies in running waters (almost exclusively in rivers), the term microinvertebrates can be replaced by zooplankton. Zooplankton are small invertebrates that live suspended in the water column and have limited ability to control their location [2]. Studies of running water macroinvertebrates vastly exceed those of zooplankton. The River Continuum project [16] recognized zooplankton as a dominant group in rivers of orders 7–10 together with benthic macroinvertebrate such as oligochaetes, bivalve gastropods, and some micro-crustaceans [1]. Studies of river zooplankton have focused on their role in food chains of fish and organic matter cycling. Although most of the zooplankton inhabit the water column of the river where they filter feed on suspended FPOM [32] the benthic forms filter FPOM from the water column and on depositional FPOM [1, 2, 24].

#### **3. Habitats of lotic invertebrates**

#### **3.1 Erosional habitats**

Erosional habitats encompass coarse sediments (boulders, cobbles, and gravel) and large wood debris in fast-flowing water (riffles and runs). Because these habitats are well oxygenated, they normally support invertebrate populations that are the most sensitive to degradation of water quality. Microbes in the organic waste can reduce dissolved oxygen levels sufficient to stress-sensitive invertebrates [33, 34]. Erosional habitats are normal features of headwater streams and mid-sized rivers but occur less frequently in large rivers. In larger rivers, erosional habitat is found primarily in sections having a significant change in grade.

**4. Functional feeding groups (FFG)**

*DOI: http://dx.doi.org/10.5772/intechopen.88283*

*Stream Invertebrate Zoology*

Investigation of lotic invertebrates has been taxonomy based since its inception [1]. In the 1980s, Robert Penna, a major authority on freshwater invertebrates of the day [2], argued that ecology of freshwater invertebrates must be based on *species*-level taxonomy (personal communication). This goal was, and still is a valid one, but then, and even now, rarely possible to achieve. There are very few groups of lotic invertebrates for which complete species inventories have been accomplished. For example, the most recent compilation of the taxa of aquatic insects of North America [1] allows identification of almost all known genera, but not species,

collected in lotic samples. The emphasis is on immature nymphs and larvae.

enter the water in order to deposit eggs are also keyed [1].

Coleoptera adults, which are aquatic, and adult females of some aquatic insects that

In 1973 and 1974, Cummins [36, 37] argued that while efforts will continue toward expanding the taxonomy of freshwater invertebrates (especially insects), it should be possible to address ecological study of lotic macroinvertebrates by employing analyses of their morphological and behavioral adaptations. The proposal was to use five functional feeding group (FFG) adaptation categories and match them to the five basic food resource categories available in varying amounts in streams and rivers [15, 16, 26, 36, 38]. In this FFG analysis, scrapers are matched with periphyton (attached nonfilamentous algae and associated material), gathering collectors with FPOM deposited on or in the sediments and filtering collectors with FPOM transported in the water column, shredders with CPOM (conditioned plant litter or live vascular plants), and predators with their prey [15, 25, 26, 39, 40]. More recently, a refinement of two more FFG matching categories has been utilized [41]. The shredder category is divided into detrital shredders (DSH) matched with CPOM conditioned plant litter or wood [42] and herbivore shredders with live vascular aquatic plant tissue [41]. Piercers are matched with filamentous algae (**Tables 1–3**) [26, 38]. Taxa that share morphological (e.g. moth parts, body structure, color pattern) and/or behavioral (e.g. movement patterns, silk netspinning, case construction) adaptations can be grouped in the same FFG. Through parallel or convergent evolution, they share features that result in the same modes of food acquisition. An example is the striking similarity between North American Heptageniidae and Brazilian Leptophlebiidae mayfly nymphs which are both scrapers [26, 38]. Also, as shown in [26], caddisfly genera in three different families (Glossosomatidae, *Glossosoma*; Helicopsychidae, *Helicopsyche*; and Uenoidae*,* Neophylax) and a beetle genus *Psephenus* scrape substrate surfaces in riffles.

A simple picture key can be used to sort macroinvertebrates collected in lotic field samples into FFG categories with an 80% or greater accuracy. This can be accomplished using structural and behavioral characters that can be readily

observed in the field on live specimens with the unaided eye or a simple hand lens. For example, case-bearing Trichoptera can be separated based on the materials used in case construction: larvae with organic cases constructed of leaf or wood pieces are detrital shredders (DSH), and those with mineral cases made of sand or fine gravel are scrapers (SC). An example of a key used to separate Ephemeroptera nymphs (lateral abdominal gills) from Plecoptera nymphs (no lateral abdominal gills), separate mayfly scraper nymphs (clinger flat body shape) from mayfly Gathering Collector nymphs (swimmer cylindrical body shape). Dragonfly and damselfly nymphs can be separated from mayfly and stonefly nymphs by an extendible

When separating live macroinvertebrates collected in a stream field sample into

functional feeding groups, individuals in different taxa that share similar

grasping labium (**Figure 1**) [15, 26, 40].

**163**

Macroinvertebrates adapted to erosional habitats (clingers) include Gastropoda (snails, e.g. Sulcospiridae, *Juga*), Ephemeroptera (mayflies, e.g. Heptageniidae), Trichoptera (stone case-bearing and net-spinning caddis), Plecoptera (stoneflies, predaceous Perlidae), and Coleoptera (Psephenidae, water pennies). The ecological tables in Berg et al. [1] identify erosional habitat adaptations by taxonomic group. Structures, such as suckers, hook, or claws of various sorts, silk that fastens down their retreats or provides anchors, and body shape and behavior that avoids the major force of the current are the main adaptations to erosional conditions [35].

#### **3.2 Depositional habitats**

Depositional habitats are drop zones where fine sediments settle out. Substrates of sand, silt, and clay are found in pools, backwaters, and along channel margins. FPOM, and in some cases. CPOM plant litter lotter [23], also accumulates in depositional habitats. These depositional habitats are dominated by sprawlers and burrowers that move across the soft substrate or are concealed beneath it [1]. Some of the Ephemeroptera sprawlers have modified first abdominal gills that cover the remaining gills to protect them from smothering by depositional silt (e.g. Caenidae, Tricorythidae) [1]. Burrowing depositional taxa include Oligochaeta, Ephemeroptera (Ephemeridae), Diptera (Chironomidae midges), and predator Odonata dragonflies (Gomphidae) [1].

#### **3.3 Rooted aquatic vascular plants**

Aquatic vascular plants occur in both erosional or depositional habitat but are more common in the depositional areas, especially in larger rivers. The macroinvertebrates associated with vascular plants feed on floating leaves such as Lepidoptera (moth larvae, e.g. Noctuidae) and Coleoptera (beetles, e.g. Chrysomelidae, *Galeracella*) and some Diptera, Chironomidae (midges). These are all herbivore shredders (HSH) that mine leaves or burrow into stems, or feed on roots, especially of *Nuphar*, where they penetrated foot tissue and extract oxygen (Coleoptera, Chrysomelidae, *Donacia*)) [1].

In the Lepidoptera ecology table [1], all genera are described as herbivore shredders (HSH) occurring in lentic (standing water) habitats. Two families (Cosmopterigidae, 180 sp., and Noctuidae, 12 sp.) are in a category "generally lentic" because they also occur in lotic systems. The emergent and floating-leaf plant beds are in backwaters, along margins and in areas of slow current of midsized and larger rivers. Some streams of orders 2 and 3 support floating-leaf plants in erosional habitats (e.g. *Valsineria*).

#### **4. Functional feeding groups (FFG)**

**3. Habitats of lotic invertebrates**

*Inland Waters - Dynamics and Ecology*

Erosional habitats encompass coarse sediments (boulders, cobbles, and gravel) and large wood debris in fast-flowing water (riffles and runs). Because these habitats are well oxygenated, they normally support invertebrate populations that are the most sensitive to degradation of water quality. Microbes in the organic waste can reduce dissolved oxygen levels sufficient to stress-sensitive invertebrates [33, 34]. Erosional habitats are normal features of headwater streams and mid-sized rivers but occur less frequently in large rivers. In larger rivers, erosional habitat is

Macroinvertebrates adapted to erosional habitats (clingers) include Gastropoda (snails, e.g. Sulcospiridae, *Juga*), Ephemeroptera (mayflies, e.g. Heptageniidae), Trichoptera (stone case-bearing and net-spinning caddis), Plecoptera (stoneflies, predaceous Perlidae), and Coleoptera (Psephenidae, water pennies). The ecological tables in Berg et al. [1] identify erosional habitat adaptations by taxonomic group. Structures, such as suckers, hook, or claws of various sorts, silk that fastens down their retreats or provides anchors, and body shape and behavior that avoids the major force of the current are the main adaptations to erosional conditions [35].

Depositional habitats are drop zones where fine sediments settle out. Substrates of sand, silt, and clay are found in pools, backwaters, and along channel margins. FPOM, and in some cases. CPOM plant litter lotter [23], also accumulates in depositional habitats. These depositional habitats are dominated by sprawlers and burrowers that move across the soft substrate or are concealed beneath it [1]. Some of the Ephemeroptera sprawlers have modified first abdominal gills that cover the remaining gills to protect them from smothering by depositional silt (e.g. Caenidae,

Tricorythidae) [1]. Burrowing depositional taxa include Oligochaeta,

more common in the depositional areas, especially in larger rivers. The

Lepidoptera (moth larvae, e.g. Noctuidae) and Coleoptera (beetles, e.g.

Ephemeroptera (Ephemeridae), Diptera (Chironomidae midges), and predator

Aquatic vascular plants occur in both erosional or depositional habitat but are

macroinvertebrates associated with vascular plants feed on floating leaves such as

Chrysomelidae, *Galeracella*) and some Diptera, Chironomidae (midges). These are all herbivore shredders (HSH) that mine leaves or burrow into stems, or feed on roots, especially of *Nuphar*, where they penetrated foot tissue and extract oxygen

In the Lepidoptera ecology table [1], all genera are described as herbivore shredders (HSH) occurring in lentic (standing water) habitats. Two families (Cosmopterigidae, 180 sp., and Noctuidae, 12 sp.) are in a category "generally lentic" because they also occur in lotic systems. The emergent and floating-leaf plant beds are in backwaters, along margins and in areas of slow current of midsized and larger rivers. Some streams of orders 2 and 3 support floating-leaf plants

found primarily in sections having a significant change in grade.

**3.1 Erosional habitats**

**3.2 Depositional habitats**

Odonata dragonflies (Gomphidae) [1].

**3.3 Rooted aquatic vascular plants**

(Coleoptera, Chrysomelidae, *Donacia*)) [1].

in erosional habitats (e.g. *Valsineria*).

**162**

Investigation of lotic invertebrates has been taxonomy based since its inception [1]. In the 1980s, Robert Penna, a major authority on freshwater invertebrates of the day [2], argued that ecology of freshwater invertebrates must be based on *species*-level taxonomy (personal communication). This goal was, and still is a valid one, but then, and even now, rarely possible to achieve. There are very few groups of lotic invertebrates for which complete species inventories have been accomplished. For example, the most recent compilation of the taxa of aquatic insects of North America [1] allows identification of almost all known genera, but not species, collected in lotic samples. The emphasis is on immature nymphs and larvae. Coleoptera adults, which are aquatic, and adult females of some aquatic insects that enter the water in order to deposit eggs are also keyed [1].

In 1973 and 1974, Cummins [36, 37] argued that while efforts will continue toward expanding the taxonomy of freshwater invertebrates (especially insects), it should be possible to address ecological study of lotic macroinvertebrates by employing analyses of their morphological and behavioral adaptations. The proposal was to use five functional feeding group (FFG) adaptation categories and match them to the five basic food resource categories available in varying amounts in streams and rivers [15, 16, 26, 36, 38]. In this FFG analysis, scrapers are matched with periphyton (attached nonfilamentous algae and associated material), gathering collectors with FPOM deposited on or in the sediments and filtering collectors with FPOM transported in the water column, shredders with CPOM (conditioned plant litter or live vascular plants), and predators with their prey [15, 25, 26, 39, 40].

More recently, a refinement of two more FFG matching categories has been utilized [41]. The shredder category is divided into detrital shredders (DSH) matched with CPOM conditioned plant litter or wood [42] and herbivore shredders with live vascular aquatic plant tissue [41]. Piercers are matched with filamentous algae (**Tables 1–3**) [26, 38]. Taxa that share morphological (e.g. moth parts, body structure, color pattern) and/or behavioral (e.g. movement patterns, silk netspinning, case construction) adaptations can be grouped in the same FFG. Through parallel or convergent evolution, they share features that result in the same modes of food acquisition. An example is the striking similarity between North American Heptageniidae and Brazilian Leptophlebiidae mayfly nymphs which are both scrapers [26, 38]. Also, as shown in [26], caddisfly genera in three different families (Glossosomatidae, *Glossosoma*; Helicopsychidae, *Helicopsyche*; and Uenoidae*,* Neophylax) and a beetle genus *Psephenus* scrape substrate surfaces in riffles.

A simple picture key can be used to sort macroinvertebrates collected in lotic field samples into FFG categories with an 80% or greater accuracy. This can be accomplished using structural and behavioral characters that can be readily observed in the field on live specimens with the unaided eye or a simple hand lens. For example, case-bearing Trichoptera can be separated based on the materials used in case construction: larvae with organic cases constructed of leaf or wood pieces are detrital shredders (DSH), and those with mineral cases made of sand or fine gravel are scrapers (SC). An example of a key used to separate Ephemeroptera nymphs (lateral abdominal gills) from Plecoptera nymphs (no lateral abdominal gills), separate mayfly scraper nymphs (clinger flat body shape) from mayfly Gathering Collector nymphs (swimmer cylindrical body shape). Dragonfly and damselfly nymphs can be separated from mayfly and stonefly nymphs by an extendible grasping labium (**Figure 1**) [15, 26, 40].

When separating live macroinvertebrates collected in a stream field sample into functional feeding groups, individuals in different taxa that share similar

#### **Figure 1.**

*A simple picture key for separating nymphal stream insects into functional feeding groups (FFG). Mayflies and stoneflies are separated by the presence or absence of lateral abdominal gills. Mayflies are separated into scrapers or gathering collectors by body shape. Stoneflies are into detrital shredders or predators by color pattern and activity level. Dragonflies and damselflies are all predators and are separated from all the other nymphs by having extensible, grasping labia (lower lips). Modified from [38].*

adaptations are enumerated together. All muffin tin with the wells is marked by FFGs (e.g. SC, GC, etc.). For example, all snails, dorsal-ventrally flattened mayfly nymphs, mineral case-bearing caddisfly larvae, and water-penny beetle larvae are sorted into the scraper (SC) category [40]. Because scrapers usually feed on surfaces in riffles, they are adapted to maintain their location; they are clingers on the substrate surface. Common North American scraper taxa found in running waters are Gastropoda snails that have a rasping radula; Ephemeroptera mayfly Heptageniidae and Ephemerellidae *Drunella* nymphs that are dorsal-ventrally flattened; Trichoptera caddisfly Limnephilidae, *Hydatophylax*, *Pycnopsyche*, Uenoidae, Glossosomatidae, and Helicopsychidae larvae; and flattened Coleoptera waterpenny beetle Psephenidae larvae (**Tables 1–4**) [1, 40]. Similar taxonomic groupings for gathering and filtering collectors, detrital shredders, herbivore shredders, or predarorsare given in **Table 4** and [40].

#### **5. Macroinvertebrates used for evaluation of lotic ecosystem condition**

The known North American aquatic insect species in genera for each order found in lotic habitats are listed in the ecological tables in Merritt et al. [43]. No lentic (standing water) genera are included in **Table 4**. The lotic genera are also assigned to FFGs in **Table 4**. Every FFG entry in **Table 4** is divided into % obligate or facultative number of species. The obligate category is defined as having a maximized % conversion of ingestion to growth. Obligate taxa are predicted to be most affected by environmental changes that alter their food resource. By contrast, the facultative forms are predicted to have flexible food requirements and to be better adapted to adjust to changes in food supplies, but the conversion of ingestion

**Taxa**

**165**

**Functional**

**Scrapers**

**Fac.**

**Ephemeroptera**

Heptageniidae Leptophlebiidae

Baetidae Ephemeridae Ephemerellidae

Isonychidae **Plecoptera**

Pteronarcyidae

Peltoperlidae

Nemouridae

Leuctridae

Capniidae Taeniopterygidae

Perlidae Perlodidae Chloroperlidae

**Trichoptera**

Glossosomatidae

Helicopsychidae

Uenoidae

0

0

0

 53.6

 53.6

 46.4

 0

 46.4

 100

 100

 71.0

 71.0

 29.0

 0

 29.0

1.4

2.1

 0

 2.1

 31.9

 0

 31.9

 0

 1.4

 28.1

 0

 28.1

22.5

 0

 22.5

 10.8 13.3

 0

 13.3

 0

 10.5

10.0

11.2

2.6

 0

 2.6

 8.3 8.3

 0

 8.3

 0

 8.3

 2.0

 13.2

 22.0

 0

 22.0

 0

 10.0

32.0

 280

 60.0

0

15.2

27.0

 11.2

 38.2

 74.5 50.0

 0

 50.0

 0 0

 100

 100 10.0

49.3

89.1

0

0

22.5

 44.1

 66.5 0

0

0

 66.0

 66.0

 70.5

 70.5

 86.7

 86.7

 100

 100

 74.8

 74.8

 0

 89.1

 11.3

 60.6

 4.4

 0

 4.4

 40.0

 59.0

 49.0

 0

 40.0

 50.0

 50.0

 43.9

 43.9

 4.0

 0

 4.9

 3.3

 18.5

 51.0

 44.0

 95.0

30.3

 0

 30.3

 49.5

 49.5

 49.5

 0

 49.5

 **Obl.**

 **Total**

 **Fac.**

 **Obl.**

 **Total**

 **Fac.**

 **Obl.**

 **Total**

 **Fac.**

 **Obl.**

 **Total**

 **Fac.**

0

 1.0

 1.0

*DOI: http://dx.doi.org/10.5772/intechopen.88283*

 **Obl.**

 **Total**

*Stream Invertebrate Zoology*

**Gathering collectors**

 **feeding group (FFG) categories**

 **Filtering collectors**

**Shredders**

**Predators**


#### *Stream Invertebrate Zoology DOI: http://dx.doi.org/10.5772/intechopen.88283*

adaptations are enumerated together. All muffin tin with the wells is marked by FFGs (e.g. SC, GC, etc.). For example, all snails, dorsal-ventrally flattened mayfly nymphs, mineral case-bearing caddisfly larvae, and water-penny beetle larvae are sorted into the scraper (SC) category [40]. Because scrapers usually feed on surfaces in riffles, they are adapted to maintain their location; they are clingers on the substrate surface. Common North American scraper taxa found in running waters

*A simple picture key for separating nymphal stream insects into functional feeding groups (FFG). Mayflies and stoneflies are separated by the presence or absence of lateral abdominal gills. Mayflies are separated into scrapers or gathering collectors by body shape. Stoneflies are into detrital shredders or predators by color pattern and activity level. Dragonflies and damselflies are all predators and are separated from all the other nymphs by*

Heptageniidae and Ephemerellidae *Drunella* nymphs that are dorsal-ventrally flattened; Trichoptera caddisfly Limnephilidae, *Hydatophylax*, *Pycnopsyche*, Uenoidae, Glossosomatidae, and Helicopsychidae larvae; and flattened Coleoptera waterpenny beetle Psephenidae larvae (**Tables 1–4**) [1, 40]. Similar taxonomic groupings for gathering and filtering collectors, detrital shredders, herbivore shredders, or

**5. Macroinvertebrates used for evaluation of lotic ecosystem condition**

The known North American aquatic insect species in genera for each order found in lotic habitats are listed in the ecological tables in Merritt et al. [43]. No lentic (standing water) genera are included in **Table 4**. The lotic genera are also assigned to FFGs in **Table 4**. Every FFG entry in **Table 4** is divided into % obligate or facultative number of species. The obligate category is defined as having a maximized % conversion of ingestion to growth. Obligate taxa are predicted to be most affected by environmental changes that alter their food resource. By contrast, the facultative forms are predicted to have flexible food requirements and to be better adapted to adjust to changes in food supplies, but the conversion of ingestion

are Gastropoda snails that have a rasping radula; Ephemeroptera mayfly

predarorsare given in **Table 4** and [40].

*having extensible, grasping labia (lower lips). Modified from [38].*

*Inland Waters - Dynamics and Ecology*

**Figure 1.**

**164**


**Taxa**

**167**

**Functional**

**Scrapers**

**Fac.**

**Lepidoptera**

Crambidae,

Noctuide

**Coleoptera**

Larvae and Adults Dytiscidae, Gyrinidae

Hydrophilidae

Hydrophilidae

0

 100

 100 100

 0

 100 100

 0

 100

Adults

**Diptera**

Tipulidae,*Tipula*

*Holorusia*

Dicranota, Pedicia,

Chironomidae

0

26.3

 *among the FFG categories representing that taxon. Fac. = facultative taxa, that is, these taxa are listed in the ecological tables in [43] as having*

 11.3

 37.6

0

 73.7

 73.7 *accompanying*

*accompanying*

 *Fac., this means that no Fac. alternatives were given in*

 *obligatory (Obl.) entry for*

 84.2

 84.2

 12.2

 0

 12.2

Chironomini

Tanytarsini

*For each taxon (column 1), the % species for each taxon is apportioned*

*several alternative FFG*  *that taxon is 0, this means there were no Fac. possibilities listed in [43]. Similarly, if there is an Obl. entry in the table and a 0 given for the* 

*entries for that taxon in [43]. If there are % entries in both columns, there are values given for both categories in [43]. When selecting the % to be used to assign the proportion of each FFG to each taxon, the clearest*

*approach would be to use only the Obl. designations* *probable Fac. species per genus. The % values in this table will be subject to some changes when [1] is published.*

*1Early instars of larvae with organic cases are obligate Detrital Shredders (DSH).*

*2Last (5th) instar of larvae with mineral cases are obligate Scrapers (SC).*

*3Because of uncertain* 

**Table 4.** *Percent species in genera of North American lotic* 

*macroinvertebrates*

 *[43].*

*Chironomidae*

 *taxonomy, the FFG percent values are based on approximate*

 *species per genus numbers [43].*

 *throughout; the most conservative*

 *approach would be to use the total % values, that is, combining those taxa that are restricted to a given FFG with the most*

*classifications.*

 *The Fac. values given in the table are the FFG that appears first (as the most likely) of the alternatives of the several presented. If the* 

*Hexatoma*

 Larvae

 **Obl.**

 **Total**

 **Fac.**

 **Obl.**

 **Total**

 **Fac.**

 **Obl.**

 **Total**

 **Fac.**

0

 100

 100 0

 100

 100

*DOI: http://dx.doi.org/10.5772/intechopen.88283*

 **Obl.**

 **Total**

 **Fac.**

 **Obl.**

 **Total**

*Stream Invertebrate Zoology*

**Gathering collectors**

 **feeding group (FFG) categories**

 **Filtering collectors**

**Shredders**

**Predators**


*Stream Invertebrate Zoology DOI: http://dx.doi.org/10.5772/intechopen.88283*

> **4.** *Percent species in genera*

 *of North American lotic* 

*macroinvertebrates*

 *[43].*

**Table**

*2Last (5th) instar of larvae with mineral cases are obligate Scrapers (SC).*

*3Because of uncertain* 

*Chironomidae*

 *taxonomy, the FFG percent values are based on approximate*

 *species per genus numbers [43].*

**Taxa**

**166**

**Functional**

**Scrapers**

**Fac.**

6.4

 4.11

10.5

 33.9 33.3

0

 22.9

 22.9

 0

 33.3

 4.1

 38.0

Limnephilidae Lepidostomatidae

Calamoceratidae

Leptoceridae Rhyacophilidae

Brachycentridae

Hydropsychidae

Philopotamidae

Psychomyiidae Polycentropidae

**Megaloptera**

Corydalidae

Sialidae **Odonata**

Anisoptera

Zygoptera **Hemiptera**

Corixidae

Nepidae Naucoridae

100

 0

 100

33.8

 20.0

 53.8

 24.6

 21.5

 46.1 42.6

 5.8

 48.4

 3.6

 0

 3.6

 7.2

0

0

0

0

0

0

 100

 100

 100

 100

 100

 100

 100

 100

 100

 100

 100

 100

 40.6

 47.8

0.8

 0

 0.8 38.8

0

 21.8

 21.8

 0

 0 0

 100

 100

 78.2

 78.2

 0

 38.8

 0

 33.8

 33.8

 **Obl.**

 **Total**

 **Fac.**

 **Obl.**

 **Total**

 **Fac.**

 **Obl.**

 **Total**

 **Fac.**

3.5

0

0

39.6

 0

 39.6

 8.3

0

 99.2

 99.2

 29.2

 37.5

 66.7

 66.7

 100

 100

*Inland Waters - Dynamics and Ecology*

 48.02

51.5

 **Obl.**

 **Total**

 **Fac.**

 **Obl.**

 **Total**

**Gathering collectors**

 **feeding group (FFG) categories**

 **Filtering collectors**

**Shredders**

**Predators**

to growth would be less efficient. Facultative taxa would be predicted to better survive environmental changes [15, 43]. When selecting the % to be used to assign the proportion of each FFG into each taxon, the least ambiguous approach would be to use only the obligate designations throughout; the most conservative approach would be to use the total % values, that is, combining those obligate taxa which are restricted to a given FFG with the most probable facultative species per genus. The % values in **Table 4** undoubtedly will be subject to some changes when [1] is published.

Using counts of numerical abundance of macroinvertebrates in field samples, ratios of the % numerical abundance of FFGs, like those in **Table 4**, can be used to calculate ratios of the FFGs. These ratios have been used as surrogates for stream and river ecosystem attributes [1, 26, 40, 41]. Because such ratios are dimensionless numbers, the resulting calculated ratios are essentially independent of sample size. For example, the FFG ratio from one riffle (coarse sediment) sample produces the same ratio as five samples, or one plant litter sample is the same as five.

A number of FFG ratios that serve as surrogates for running water ecosystem attributes are summarized in **Table 5**. Thresholds for evaluating the ratios are also proposed ([1], Table 6E). The ecosystem attributes can be measured directly, but this usually requires significant equipment, time, and direct tending by researchers. In addition, the actual measurements represent only a fraction of the temporal and special scales at which the processes occur. By contrast, the macroinvertebrates continuously monitor ecosystem conditions over their life stages in the water, at least weeks and usually annual or semiannual periods.

Arguably, the most all-encompassing and informative ratio is gross primary production (P) compared to community respiration. The P/R ratio also reflects the relative dominance of autotrophy (energy source within the stream or river relative to energy input from outside the aquatic ecosystem) (**Table 5**) [44–46]. The surrogate macroinvertebrate P/R ratio is all FFGs that depend on autochthonous primary production (algae and vascular plants) compared to all the FFGs that depend on FPOM and CPOM organic matter. That is, scrapers + herbivore shredders + algal cell piercers to detrital shredders + gathering collectors + filtering collectors (**Table 5**) [1, 10, 41, 47]. The P/R ratio that corresponds to a directly measured P/R, using closed, recirculating chambers that monitor dissolved oxygen, of P/R > 1.0 is a macroinvertebrate P/R > 0.75 (**Table 5**) [19, 44, 45]. The other surrogate ratios described compare detrital shredders available CPOM storage (Detrital Shredder index), relative abundance of filtering collectors to FPOM in transport (Filtering Collector Index), macroinvertebrates that require stable attachment or clinging sites compared to substrate stability, and predator abundance relative to prey available (Predator Index) (**Table 5**).

The ecological tables in [1, 43] also include US Environmental Protection Agency values for macroinvertebrate susceptibility/resistance that are indicators of pollution. As a general rule, the EPT Index will indicate the vulnerability of macroinvertebrates to stream and river water quality degradation. This index compares the abundance of Ephemeroptera (mayflies) + Plecoptera (stoneflies) + Trichoptera (caddisflies) to the rest of the macroinvertebrate fauna; the more dominant the EPT, the less polluted the stream or river is rated [33, 48].

Organic pollution reduces dissolved oxygen (DO) levels in freshwater due to the large oxygen demand by microbial respiration [33]. Significant reduction in DO is a major stressor for aerobic (DO requiring) invertebrates. This includes those with gills or some with cutaneous respiration: Mollusca, Crustacea, Ephemeroptera, Plecoptera, Odonata, Trichoptera, Megaloptera, some Lepidoptera, some Coleoptera larvae, and some Diptera. Oligochaeta and some Chironomidae have biochemical adaptations that allow them to tolerate low DO levels [1].

However, there are stream and river macroinvertebrates that are adapted to breathe atmospheric oxygen (AO) by returning to the water surface to obtain air, such as some Diptera (the best known example being Culicidae mosquitoes). Others like some Coleoptera adults trap air in body surface hairs or under elytra that they carry under the surface and from which they extract DA. An index that would

*Functional feeding group (FFG) ratios as surrogates for running water lotic ecosystem attributes and proposed*

**FFG ratios Symbols Proposed**

SC + HSH to DSH + GC + FC

SC + FC + HSH to DSH + GC

P to SC + DSH + HSH FC + FC

DSH to GC + FC CPOM/FPOM

Scrapers + herbivore shredders to detrital shredders + gathering collectors + filtering collectors

Detrital shredders to gathering collectors + filtering collectors

Filtering collectors to gathering collectors

Scrapers + filtering collectors + herbivore shredders to detrital shredders + gathering collectors

Predators to scrapers + detrital shredders + herbivore shedders + filtering collectors + gathering collectors

**thresholds**

>0.5 (fall– winter); CPOM/FPOM >0.25 (spring– summer)

FC to GC FC/GC > 0.5 FC > 0.5 favors

Stable substrates/ unstable substrates >0.5

Predators/all other FFGs present =0.10–0.15

**Descriptions**

corresponds to P/R > 1 when primary production and community respiration are measured directly

The CPOM/FPOM >0.5 during fall– winter when fast processed deciduous plant litter enters streams; > 0.25 in spring–summer when slower processed needles and wood remain in streams

filtering collectors capturing FPOM in transport; impaired lotic systems usually have values much higher than 9.5

Stable substrates (bedrock, boulders and cobbles, large wood, and rooted plants) provide attachment and clinger sites greater than unstable fine sands and clay. Channel disturbance can reduce stable substrates or flush out fines

The abundance of predators between 10% and 15% of the lotic macroinvertebrate community indicates sufficient prey (turnover to support predators)

P/R > 0.75 A P/R > 0.75

**Lotic ecosystem attributes**

*Stream Invertebrate Zoology*

*DOI: http://dx.doi.org/10.5772/intechopen.88283*

Autotrophic to heterotrophic Index (P/R, primary production/ community respiration)

Shredder Index (CPOM/FPOM) [27, 28]

Filtering collector Index (suspended FPOM to deposited FPOM)

Substrate stability Index (stable coarse substrates to unstable fine sediments)

Predator Index (top-down to bottom-up control

macroinvertebrate communities)

of

**Table 5.**

*thresholds.*

**169**

#### *Stream Invertebrate Zoology DOI: http://dx.doi.org/10.5772/intechopen.88283*

to growth would be less efficient. Facultative taxa would be predicted to better survive environmental changes [15, 43]. When selecting the % to be used to assign the proportion of each FFG into each taxon, the least ambiguous approach would be to use only the obligate designations throughout; the most conservative approach would be to use the total % values, that is, combining those obligate taxa which are restricted to a given FFG with the most probable facultative species per genus. The % values in **Table 4** undoubtedly will be subject to some changes when [1] is

Using counts of numerical abundance of macroinvertebrates in field samples, ratios of the % numerical abundance of FFGs, like those in **Table 4**, can be used to calculate ratios of the FFGs. These ratios have been used as surrogates for stream and river ecosystem attributes [1, 26, 40, 41]. Because such ratios are dimensionless numbers, the resulting calculated ratios are essentially independent of sample size. For example, the FFG ratio from one riffle (coarse sediment) sample produces the

A number of FFG ratios that serve as surrogates for running water ecosystem attributes are summarized in **Table 5**. Thresholds for evaluating the ratios are also proposed ([1], Table 6E). The ecosystem attributes can be measured directly, but this usually requires significant equipment, time, and direct tending by researchers. In addition, the actual measurements represent only a fraction of the temporal and special scales at which the processes occur. By contrast, the macroinvertebrates continuously monitor ecosystem conditions over their life stages in the water, at

Arguably, the most all-encompassing and informative ratio is gross primary production (P) compared to community respiration. The P/R ratio also reflects the relative dominance of autotrophy (energy source within the stream or river relative to energy input from outside the aquatic ecosystem) (**Table 5**) [44–46]. The surrogate macroinvertebrate P/R ratio is all FFGs that depend on autochthonous primary production (algae and vascular plants) compared to all the FFGs that depend on FPOM and CPOM organic matter. That is, scrapers + herbivore shredders + algal cell piercers to detrital shredders + gathering collectors + filtering collectors

(**Table 5**) [1, 10, 41, 47]. The P/R ratio that corresponds to a directly measured P/R, using closed, recirculating chambers that monitor dissolved oxygen, of P/R > 1.0 is a macroinvertebrate P/R > 0.75 (**Table 5**) [19, 44, 45]. The other surrogate ratios described compare detrital shredders available CPOM storage (Detrital Shredder index), relative abundance of filtering collectors to FPOM in transport (Filtering Collector Index), macroinvertebrates that require stable attachment or clinging sites compared to substrate stability, and predator abundance relative to prey avail-

The ecological tables in [1, 43] also include US Environmental Protection Agency values for macroinvertebrate susceptibility/resistance that are indicators of pollution. As a general rule, the EPT Index will indicate the vulnerability of macroinvertebrates to stream and river water quality degradation. This index compares the abundance of Ephemeroptera (mayflies) + Plecoptera (stoneflies) + Trichoptera (caddisflies) to the rest of the macroinvertebrate fauna; the more dominant the

Organic pollution reduces dissolved oxygen (DO) levels in freshwater due to the large oxygen demand by microbial respiration [33]. Significant reduction in DO is a major stressor for aerobic (DO requiring) invertebrates. This includes those with gills or some with cutaneous respiration: Mollusca, Crustacea, Ephemeroptera, Plecoptera, Odonata, Trichoptera, Megaloptera, some Lepidoptera, some Coleoptera larvae, and some Diptera. Oligochaeta and some Chironomidae have biochem-

same ratio as five samples, or one plant litter sample is the same as five.

least weeks and usually annual or semiannual periods.

EPT, the less polluted the stream or river is rated [33, 48].

ical adaptations that allow them to tolerate low DO levels [1].

able (Predator Index) (**Table 5**).

**168**

published.

*Inland Waters - Dynamics and Ecology*


#### **Table 5.**

*Functional feeding group (FFG) ratios as surrogates for running water lotic ecosystem attributes and proposed thresholds.*

However, there are stream and river macroinvertebrates that are adapted to breathe atmospheric oxygen (AO) by returning to the water surface to obtain air, such as some Diptera (the best known example being Culicidae mosquitoes). Others like some Coleoptera adults trap air in body surface hairs or under elytra that they carry under the surface and from which they extract DA. An index that would

predict poor ability of lotic macroinvertebrate taxa to survive under declining DO conditions would be taxa with gills as a proportion of those taxa with adaptations, structural, behavioral, and/or biochemical, that allows them to breath AO.

and chemical grab samples or even recording electrodes. Unlike algal cells and microbes, macroinvertebrates can be observed with the naked eye and a simple hand lens. They are far less migratory than fish that respond to an environmental stressor by leaving. Because running water macroinvertebrates are ubiquitous, eas-

ily collected, and observed and can be classified into meaningful categories

this promising track.

*Stream Invertebrate Zoology*

*DOI: http://dx.doi.org/10.5772/intechopen.88283*

**Author details**

**171**

Kenneth W. Cummins

\*Address all correspondence to: kc8161@gmail.com

provided the original work is properly cited.

thatchronical the condition of freshwater resources, they are the perfect vehicle for use by basically and easily trained local volunteers. This yields a potential army of environmental stewards who can enlist our very best freshwater monitors—the invertebrates. Both conventional morphological taxonomy and DNA barcoding will undoubtedly continue to lead to ever-better answers to what is it (classification), but this is only the initial step to answering the ultimate question—what does it do (function)? So, students, researchers, and armature naturalists, let us continue on

Cooperative Fisheries Research Unit, Humboldt State University, Arcata, CA, USA

© 2019 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

The most widely used method to obtain qualitative samples in streams and shallow rivers is the D-frame dip net. The net usually has a 1.0 or 0.5 mm mesh size. However, nets with a 0.25 mesh are recommended to retain midges (Chironomidae and Ceratopogonidae) and early stages or instars of macroinvertebrate taxa. The samples collected with the D-frame net can be considered semiquantitative when fixed time (e.g. 30 s) sampling is employed. This method has been used to compare stream/river reaches, stream/river habitats, seasons, etc. D-frame net sampling has also been used to collect composite samples, that is, an effort to collect widely from all habitats in a stream or river reach [41, 47]. In such complete reach survey samples, it is useful to keep the major habitat samples (riffles, pools, CPOM plant litter accumulations, large wood) separate and compute a composite value by calculating a total value by combining the data after the samples have been processed. This method is even more useful if a percent of the stream/river bottom in the sample reach is covered by each habitat type. This is particularly helpful if the derived FFG ratios described above are to be used as surrogates for lotic ecosystem attributes. If the same person is used to collect the D-frame samples, the results are more comparable; if that same person is also used to collect quantitative Surber net samples, the variances are similar. Ratios and proposed threshold values are presented in **Table 5.**

#### **6. Quantitative sampling**

The two most commonly used quantitative devices used for sampling stream macroinvertebrates are the Surber and Hess samplers [3, 49]. Both of these confine an area of stream bottom and rely on the stream current to transport material disturbed by the hand into attached collection nets. The Surber net has a metal frame that delineates the bottom area to be collected with an erect frame at the back that holds the collection net. The net is washed into a sorting tray (usually a white enamel pan) for partial or complete processing in the field, or the contents are rinsed directly into a bottle and preserved in 70% ETOH. The Hess sampler is a cylinder that defines the bottom area sampled. It has mesh side panels and a collection net to retain material disturbed from the bottom similar to the Surber net. With both collection devices, the sample collected is better handled if cobbles are removed individually and scrubbed into the net and discarded. As with qualitative sampling, the net mesh size is a significant issue. Both Surber and Hess samplers are available with 1 or 0.5 mm nesh. However, as stated previously, a 0.25 -mm mesh is better because it will retain early instars and smaller species are lost with the coarser mesh sizes. If the samples are to be sorted for FFGs, it is highly recommended that this be done with live specimens following collection.

#### **7. Concluding remarks**

In the past and present, and predictably in the future, abundance and composition of stream and river invertebrate communities have been, and will be, the primary measuring biological tool used to evaluate ecosystem condition and to predict environmental change and vulnerability. These animals, because they continuously monitor the stream and river environment throughout their aquatic life, can provide better insight than the spatially and temporally and limited physical

#### *Stream Invertebrate Zoology DOI: http://dx.doi.org/10.5772/intechopen.88283*

predict poor ability of lotic macroinvertebrate taxa to survive under declining DO conditions would be taxa with gills as a proportion of those taxa with adaptations,

structural, behavioral, and/or biochemical, that allows them to breath AO. The most widely used method to obtain qualitative samples in streams and shallow rivers is the D-frame dip net. The net usually has a 1.0 or 0.5 mm mesh size. However, nets with a 0.25 mesh are recommended to retain midges (Chironomidae and Ceratopogonidae) and early stages or instars of macroinvertebrate taxa. The samples collected with the D-frame net can be considered semiquantitative when fixed time (e.g. 30 s) sampling is employed. This method has been used to compare stream/river reaches, stream/river habitats, seasons, etc. D-frame net sampling has also been used to collect composite samples, that is, an effort to collect widely from all habitats in a stream or river reach [41, 47]. In such complete reach survey samples, it is useful to keep the major habitat samples (riffles, pools, CPOM plant litter accumulations, large wood) separate and compute a composite value by calculating a total value by combining the data after the samples have been processed. This method is even more useful if a percent of the stream/river bottom in the sample reach is covered by each habitat type. This is particularly helpful if the derived FFG ratios described above are to be used as surrogates for lotic ecosystem attributes. If the same person is used to collect the D-frame samples, the results are more comparable; if that same person is also used to collect quantitative Surber net

samples, the variances are similar. Ratios and proposed threshold values are

The two most commonly used quantitative devices used for sampling stream macroinvertebrates are the Surber and Hess samplers [3, 49]. Both of these confine an area of stream bottom and rely on the stream current to transport material disturbed by the hand into attached collection nets. The Surber net has a metal frame that delineates the bottom area to be collected with an erect frame at the back that holds the collection net. The net is washed into a sorting tray (usually a white enamel pan) for partial or complete processing in the field, or the contents are rinsed directly into a bottle and preserved in 70% ETOH. The Hess sampler is a cylinder that defines the bottom area sampled. It has mesh side panels and a collection net to retain material disturbed from the bottom similar to the Surber net. With both collection devices, the sample collected is better handled if cobbles are removed individually and scrubbed into the net and discarded. As with qualitative sampling, the net mesh size is a significant issue. Both Surber and Hess samplers are available with 1 or 0.5 mm nesh. However, as stated previously, a 0.25 -mm mesh is better because it will retain early instars and smaller species are lost with the coarser mesh sizes. If the samples are to be sorted for FFGs, it is highly recommended that

In the past and present, and predictably in the future, abundance and composi-

tion of stream and river invertebrate communities have been, and will be, the primary measuring biological tool used to evaluate ecosystem condition and to predict environmental change and vulnerability. These animals, because they continuously monitor the stream and river environment throughout their aquatic life, can provide better insight than the spatially and temporally and limited physical

presented in **Table 5.**

**6. Quantitative sampling**

*Inland Waters - Dynamics and Ecology*

**7. Concluding remarks**

**170**

this be done with live specimens following collection.

and chemical grab samples or even recording electrodes. Unlike algal cells and microbes, macroinvertebrates can be observed with the naked eye and a simple hand lens. They are far less migratory than fish that respond to an environmental stressor by leaving. Because running water macroinvertebrates are ubiquitous, easily collected, and observed and can be classified into meaningful categories thatchronical the condition of freshwater resources, they are the perfect vehicle for use by basically and easily trained local volunteers. This yields a potential army of environmental stewards who can enlist our very best freshwater monitors—the invertebrates. Both conventional morphological taxonomy and DNA barcoding will undoubtedly continue to lead to ever-better answers to what is it (classification), but this is only the initial step to answering the ultimate question—what does it do (function)? So, students, researchers, and armature naturalists, let us continue on this promising track.

### **Author details**

Kenneth W. Cummins Cooperative Fisheries Research Unit, Humboldt State University, Arcata, CA, USA

\*Address all correspondence to: kc8161@gmail.com

© 2019 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

#### **References**

[1] Berg MB, Merritt RW, Cummins KW, editors. An Introduction to the Aquatic Insects of North America. Dubuque, IA: Kendall/Hunt; 2019

[2] Smith DG. Pennak's Freshwater Invertebrates of the United States: Porifera to Crustacea. New York: Wiley; 2001

[3] Hauer FR, Resh VH. Macroinvertebrates. In: Hauer FR, Lamberti GA, editors. Methods in Stream Ecology. Volume 1: Ecosystem Structure. London: Academic Press/ Elsevier; 2017. pp. 297-319

[4] Shelford VE. An experimental study of the behavior agreement among animals of an animal community. The Biological Bulletin. 1914;**20**:294-315

[5] Shelford VE, Edy S. Methods in the study of stream communities. Ecology. 1929;**10**:381-392

[6] Moon HP. Aspects of the ecology of aquatic insects. Transactions of the British Entomological and Natural History Society. 1939;**6**:39-49

[7] Cummins KW. What is a river: Zoological description. In: Ogelsby RT, Calson CA, NcCann JA, editors. River Ecology and Man. New York: Academic Press; 1972. pp. 33-52. 465p

[8] Wessenberg-Lund C. Biologie der Susswasser Insecten. Copenhagen, Denmark: Gylden & Springer; 1943. 682p

[9] Allen RH. The Horokiwi stream. New Zealand Marine Department Fisheries Bulletin. 1951;**101**:1-231

[10] Cummins KW, Merritt RW, Berg MB. Ecology and distribution of aquatic insects. In: Berg MB, Merritt RW, Cummins KW, editors. An Introduction to the Aquatic Insects of North America. Dubuque, IA: Kendall/Hunt; 2019

Sandusky River Basin Symposium on International Joint Communications, International GP. Great Lakes Pollution from Land Use Activities. Washington, DC: U. S. Government Printing Office;

*DOI: http://dx.doi.org/10.5772/intechopen.88283*

[25] Benfield EF, Fritz KM. Leaf litter breakdown. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London:

[26] Cummins KW. Functional analysis of stream macroinvertebrates. In: Limnology—Some New Aspects of Inland Water Ecology. Rijeka, Croatia:

[27] Grubbs SA, Cummins KW. Linkages between Riparian forest composition and shredder voltinism. Archiv für Hydrobiologie. 1996;**137**:49-58

[28] Wessell KJ, Merritt RW, Wilhelm

[29] Jackson JK, Batzed DP, Resh VH. Chapter 3: Sampling aquatic insects: Collection devices, statistical

consideration, and rearing procedures. In: Berg MB, Merritt RW, Cummins KW, editors. An introduction to the aquatic insects of North America. 5th ed. Dubuque, IA, USA: Kendall/Hunt

[30] Minshall GW, Peterson RC, Bott TL, Cushing CE, Cummins KW, Vannote RL, et al. Stream ecosystem dynamics of the Salmon River, Idaho: An 8th order system. JNABS. 1992;**11**:

[31] Rugenski AT, Minshall GW, Hauer FR. Riparian processes and interactions. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic

[32] Schailendra S, Siddique A, Singh K, Chouhan M, Vyas A, Solnki C, et al.

Press/Elsevier; 2017. pp. 83-111

JGO, Allan JD, Cummins KW, Uzarski DG. Biological evaluation of Michigan's non-wadeable rivers using macroinvertebrates. Aquatic Ecosystem Health & Management.

2005;**11**:35-351

Publ. Co.; 2019

111-137

Academic Press/Elsevier; 2017.

IntechOpen; 2018. DOI: 105772intechopen.79913

pp. 71-82

[19] Cummins KW, Klug MJ, Ward GM, Spengler GL, Speaker RW, Ovink RW, et al. Trends in particulate organic matter fluxes, community processes, and and macroinvertebrate functional groups, along a Great Lakes drainage basin river continuum. Verhandlungen des Internationalen Verein Limnologie.

[20] Junk WJ, Bayley PB, Sparks RE. The flood pulse concept in river flood-plain

[21] Ward JV, Stanford JA. Ecological connectivity in alluvial river ecosystems and its disruption by flow regulation. Regulated Rivers. 1995;**11**:105-119

[22] Marchant R. The ecology of *Gammarus* running water. In: Lock MA, Williams DD, editors.

New York: Plenum Press; 1981.

Press/Elsevier; 2017. pp. 55-69

Elsevier; 2017. pp. 47-53

**173**

Perspectives in Running Water Ecology.

[23] Lamberti GA, Entrekin SA, Tiegs SD. Coarse particulate organic matter: Storage, transport, and retention. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic

[24] Hutchens, Wallace JB, Grubough JW. Transport and storage of fine particulate organic matter. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic Press/

systems. In: Proceedings of the International Large River Symposium. Canadian Special Publication of Fisheries and Aquatic Sciences; 1989.

1976. pp. 277-293. 475 p

*Stream Invertebrate Zoology*

1981;**21**:841-849

p. 106

pp. 225-249

[11] Ball SL, Armstrong KF. DNA barcodes for insect pest identification: A test case with Tussock moths (Lepidoptera; Lymantridae). Canadian Journal of Forest Research. 2006;**36**: 337-350

[12] Cordero RD, Sanchez-Damirez S, Currie DC. DNA barcoding of aquatic insects reveals unforeseen diversity and recurrent population divergence patens through broad scale sampling in northern Canada. Polar Biology. 2017; **40**:1687-1695

[13] Hershey AE, Northinton JC, Finley C, Peterson BJ. Stable isotopes in stream food webs. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic Press/Elsevier; 2017. pp. 3-20

[14] Peterson BJ, Fry B. Stable isotopes in ecosystem studies. Annual Review of Ecology and Systematics. 2004;**16**: 273-320

[15] Cummins KW, Klug MJ. Feeding ecology of stream invertebrates. Annual Review of Ecology and Systematics. 1979;**10**:147-172

[16] Vannote RL, Minshall GW, Cummins KW, Sedell JR, Cushing CE. The river continuum concept. Canadian Journal of Fisheries and Aquatic Sciences. 1980;**37**:130-137

[17] Strahler AN. Hyposometric (areaaltitude) analysis of erosional topography. Americans Bulletin. 1952; **63**:1117-1142

[18] Cummins KW. The ecology of running waters: Theory and practice. In: Baker DB, Jackson WB, Prater BL, editors. The Ecology of Running Waters. In: 1975 Proceedings of the

#### *Stream Invertebrate Zoology DOI: http://dx.doi.org/10.5772/intechopen.88283*

Sandusky River Basin Symposium on International Joint Communications, International GP. Great Lakes Pollution from Land Use Activities. Washington, DC: U. S. Government Printing Office; 1976. pp. 277-293. 475 p

**References**

2001

[1] Berg MB, Merritt RW, Cummins KW, editors. An Introduction to the Aquatic Insects of North America. Dubuque, IA: Kendall/Hunt; 2019

*Inland Waters - Dynamics and Ecology*

to the Aquatic Insects of North America. Dubuque, IA: Kendall/Hunt; 2019

(Lepidoptera; Lymantridae). Canadian Journal of Forest Research. 2006;**36**:

[12] Cordero RD, Sanchez-Damirez S, Currie DC. DNA barcoding of aquatic insects reveals unforeseen diversity and recurrent population divergence patens through broad scale sampling in northern Canada. Polar Biology. 2017;

[13] Hershey AE, Northinton JC, Finley C, Peterson BJ. Stable isotopes in stream food webs. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic Press/Elsevier; 2017. pp. 3-20

[14] Peterson BJ, Fry B. Stable isotopes in ecosystem studies. Annual Review of Ecology and Systematics. 2004;**16**:

[15] Cummins KW, Klug MJ. Feeding ecology of stream invertebrates. Annual Review of Ecology and Systematics.

[16] Vannote RL, Minshall GW, Cummins KW, Sedell JR, Cushing CE. The river continuum concept. Canadian

Journal of Fisheries and Aquatic Sciences. 1980;**37**:130-137

altitude) analysis of erosional

[17] Strahler AN. Hyposometric (area-

topography. Americans Bulletin. 1952;

[18] Cummins KW. The ecology of running waters: Theory and practice. In: Baker DB, Jackson WB, Prater BL, editors. The Ecology of Running Waters. In: 1975 Proceedings of the

[11] Ball SL, Armstrong KF. DNA barcodes for insect pest identification: A

test case with Tussock moths

337-350

**40**:1687-1695

273-320

1979;**10**:147-172

**63**:1117-1142

[2] Smith DG. Pennak's Freshwater Invertebrates of the United States: Porifera to Crustacea. New York: Wiley;

Macroinvertebrates. In: Hauer FR, Lamberti GA, editors. Methods in Stream Ecology. Volume 1: Ecosystem Structure. London: Academic Press/

[4] Shelford VE. An experimental study of the behavior agreement among animals of an animal community. The Biological Bulletin. 1914;**20**:294-315

[5] Shelford VE, Edy S. Methods in the study of stream communities. Ecology.

[6] Moon HP. Aspects of the ecology of aquatic insects. Transactions of the British Entomological and Natural History Society. 1939;**6**:39-49

[7] Cummins KW. What is a river: Zoological description. In: Ogelsby RT, Calson CA, NcCann JA, editors. River Ecology and Man. New York: Academic

[8] Wessenberg-Lund C. Biologie der Susswasser Insecten. Copenhagen, Denmark: Gylden & Springer; 1943.

[9] Allen RH. The Horokiwi stream. New Zealand Marine Department Fisheries

[10] Cummins KW, Merritt RW, Berg MB. Ecology and distribution of aquatic insects. In: Berg MB, Merritt RW, Cummins KW, editors. An Introduction

Press; 1972. pp. 33-52. 465p

Bulletin. 1951;**101**:1-231

682p

**172**

[3] Hauer FR, Resh VH.

Elsevier; 2017. pp. 297-319

1929;**10**:381-392

[19] Cummins KW, Klug MJ, Ward GM, Spengler GL, Speaker RW, Ovink RW, et al. Trends in particulate organic matter fluxes, community processes, and and macroinvertebrate functional groups, along a Great Lakes drainage basin river continuum. Verhandlungen des Internationalen Verein Limnologie. 1981;**21**:841-849

[20] Junk WJ, Bayley PB, Sparks RE. The flood pulse concept in river flood-plain systems. In: Proceedings of the International Large River Symposium. Canadian Special Publication of Fisheries and Aquatic Sciences; 1989. p. 106

[21] Ward JV, Stanford JA. Ecological connectivity in alluvial river ecosystems and its disruption by flow regulation. Regulated Rivers. 1995;**11**:105-119

[22] Marchant R. The ecology of *Gammarus* running water. In: Lock MA, Williams DD, editors. Perspectives in Running Water Ecology. New York: Plenum Press; 1981. pp. 225-249

[23] Lamberti GA, Entrekin SA, Tiegs SD. Coarse particulate organic matter: Storage, transport, and retention. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic Press/Elsevier; 2017. pp. 55-69

[24] Hutchens, Wallace JB, Grubough JW. Transport and storage of fine particulate organic matter. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic Press/ Elsevier; 2017. pp. 47-53

[25] Benfield EF, Fritz KM. Leaf litter breakdown. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic Press/Elsevier; 2017. pp. 71-82

[26] Cummins KW. Functional analysis of stream macroinvertebrates. In: Limnology—Some New Aspects of Inland Water Ecology. Rijeka, Croatia: IntechOpen; 2018. DOI: 105772intechopen.79913

[27] Grubbs SA, Cummins KW. Linkages between Riparian forest composition and shredder voltinism. Archiv für Hydrobiologie. 1996;**137**:49-58

[28] Wessell KJ, Merritt RW, Wilhelm JGO, Allan JD, Cummins KW, Uzarski DG. Biological evaluation of Michigan's non-wadeable rivers using macroinvertebrates. Aquatic Ecosystem Health & Management. 2005;**11**:35-351

[29] Jackson JK, Batzed DP, Resh VH. Chapter 3: Sampling aquatic insects: Collection devices, statistical consideration, and rearing procedures. In: Berg MB, Merritt RW, Cummins KW, editors. An introduction to the aquatic insects of North America. 5th ed. Dubuque, IA, USA: Kendall/Hunt Publ. Co.; 2019

[30] Minshall GW, Peterson RC, Bott TL, Cushing CE, Cummins KW, Vannote RL, et al. Stream ecosystem dynamics of the Salmon River, Idaho: An 8th order system. JNABS. 1992;**11**: 111-137

[31] Rugenski AT, Minshall GW, Hauer FR. Riparian processes and interactions. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic Press/Elsevier; 2017. pp. 83-111

[32] Schailendra S, Siddique A, Singh K, Chouhan M, Vyas A, Solnki C, et al.

Population dynamics and seasonal abundance of zooplankton community in Narmada River (India). Research. 2010;**2**:1-9

[33] Karr JR, Chu EW. Biological monitoring: Essential foundation for ecological assessment. Human and Ecological Risk Assessment. 1997;**3L**: 993-1004

[34] Novak MA, Bode RW. Percent model affinity: A new measure of macroinvertebrate community composition. JNABS;**11**:80-85

[35] Newberry R, Wand Bates DJ. Dynamics of flowing water. In: Hauer FR, Lamberti GA, editors. Methods in Stream Ecology. Volume 1: Ecosystem Structure. London: Academic Press/ Elsevier; 2017. pp. 71-87

[36] Cummins KW. Structure and function of stream ecosystems. BioScience. 1974;**24**:631-641

[37] Cummins KW. Trophic relations in aquatic insects. Annual Review of Entomology. 1973;**18**:183-206

[38] Cummins KW. Combining taxonomy and function in the study of stream macroinvertebrates. Journal of Limnology. 2016;**75**:235-241

[39] Cummins KW, Spengler GL. Stream ecosystems. Americans Bulletin. 1978; **100**:1-9

[40] Merritt RW, Cummins KW, Berg MB. Trophic relationships of macroinvertebrates. In: Hauer FR, Lamberti GA, editors. Methods in Stream Ecology. Volume 1: Ecosystem Structure. London: Academic Press/ Elsevier; 2017. pp. 413-434

[41] Mattson RA, Cummins KW, Merritt RW, McIntosh M, Campbell E, Berg MB, et al. Hydrological monitoring of benthic invertebrate communities of the marsh habitats in the upper and middle St. Johns River. Florida Scientist. 2014; **77**:144-161

[49] Jackson JK, Batzer DP, Resh VH. Sampling aquatic insects: Collection devices, statistical consideration, and rearing procedure. In: Berg MB, Merritt

*DOI: http://dx.doi.org/10.5772/intechopen.88283*

RW, Cummins KW, editors. An Introduction to the Aquatic Insects of

North America. Dubuque, IA:

Kendall/Hunt; 2019

*Stream Invertebrate Zoology*

**175**

[42] Gregory SV, Gunell A, Piegay H, Boyer K. Dynamics of wood. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic Press/Elsevier; 2017. pp. 114-126

[43] Merritt RW, Cummins KW, Berg MB, editors. An Introduction to the Aquatic Insects of North America. Dubuque, IA: Kendall/Hunt; 2008. 1158p

[44] King DK, Cummins KW. Autotrophic-heterotrophic community metabolism relationships of a woodland stream. Journal of Freshwater Ecology. 1989;**5**:205-218

[45] King DK, Cummins KW. Factors affecting autotrophic-heterotrophic community metabolism relationships of a woodland stream. Journal of Freshwater Ecology. 1989;**5**:219-230

[46] King DK, Cummins KW. Estimates of detrital and epilithon community metabolism from particle-sized riffle sediments of a woodland stream. Journal of Freshwater Ecology. 1989;**5**: 231-246

[47] Merritt RW, Cummins KW, Berg MB, Novak JA, Higgins MJ, Wessell KJ, et al. Development and application of a macroinvertebrate functional groups approach in the bioassessment of remnant oxbows in the Caloosahatchee River, Southwest Florida. JNABS. 2002; **21**:290-310

[48] Carter JL, Resh VH, Hannaford MJ. Macroinvertebrates as biotic indicators of environmental quality. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic Press/ Elsevier; 2017. pp. 293-318

*Stream Invertebrate Zoology DOI: http://dx.doi.org/10.5772/intechopen.88283*

Population dynamics and seasonal abundance of zooplankton community in Narmada River (India). Research.

*Inland Waters - Dynamics and Ecology*

marsh habitats in the upper and middle St. Johns River. Florida Scientist. 2014;

[42] Gregory SV, Gunell A, Piegay H, Boyer K. Dynamics of wood. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic

Press/Elsevier; 2017. pp. 114-126

[44] King DK, Cummins KW.

[43] Merritt RW, Cummins KW, Berg MB, editors. An Introduction to the Aquatic Insects of North America. Dubuque, IA: Kendall/Hunt; 2008.

Autotrophic-heterotrophic community metabolism relationships of a woodland stream. Journal of Freshwater Ecology.

[45] King DK, Cummins KW. Factors affecting autotrophic-heterotrophic community metabolism relationships of

[46] King DK, Cummins KW. Estimates of detrital and epilithon community metabolism from particle-sized riffle sediments of a woodland stream. Journal of Freshwater Ecology. 1989;**5**:

[47] Merritt RW, Cummins KW, Berg MB, Novak JA, Higgins MJ, Wessell KJ, et al. Development and application of a macroinvertebrate functional groups approach in the bioassessment of remnant oxbows in the Caloosahatchee River, Southwest Florida. JNABS. 2002;

[48] Carter JL, Resh VH, Hannaford MJ. Macroinvertebrates as biotic indicators of environmental quality. In: Lamberti GA, Hauer FR, editors. Methods in Stream Ecology. Volume 2: Ecosystem Function. London: Academic Press/

Elsevier; 2017. pp. 293-318

a woodland stream. Journal of Freshwater Ecology. 1989;**5**:219-230

**77**:144-161

1158p

1989;**5**:205-218

231-246

**21**:290-310

[33] Karr JR, Chu EW. Biological monitoring: Essential foundation for ecological assessment. Human and Ecological Risk Assessment. 1997;**3L**:

[34] Novak MA, Bode RW. Percent model affinity: A new measure of macroinvertebrate community composition. JNABS;**11**:80-85

[35] Newberry R, Wand Bates DJ. Dynamics of flowing water. In: Hauer FR, Lamberti GA, editors. Methods in Stream Ecology. Volume 1: Ecosystem Structure. London: Academic Press/

[36] Cummins KW. Structure and function of stream ecosystems. BioScience. 1974;**24**:631-641

[38] Cummins KW. Combining

Limnology. 2016;**75**:235-241

MB. Trophic relationships of macroinvertebrates. In: Hauer FR, Lamberti GA, editors. Methods in Stream Ecology. Volume 1: Ecosystem Structure. London: Academic Press/

Elsevier; 2017. pp. 413-434

**100**:1-9

**174**

[37] Cummins KW. Trophic relations in aquatic insects. Annual Review of Entomology. 1973;**18**:183-206

taxonomy and function in the study of stream macroinvertebrates. Journal of

[39] Cummins KW, Spengler GL. Stream ecosystems. Americans Bulletin. 1978;

[40] Merritt RW, Cummins KW, Berg

[41] Mattson RA, Cummins KW, Merritt RW, McIntosh M, Campbell E, Berg MB, et al. Hydrological monitoring of benthic invertebrate communities of the

Elsevier; 2017. pp. 71-87

2010;**2**:1-9

993-1004

[49] Jackson JK, Batzer DP, Resh VH. Sampling aquatic insects: Collection devices, statistical consideration, and rearing procedure. In: Berg MB, Merritt RW, Cummins KW, editors. An Introduction to the Aquatic Insects of North America. Dubuque, IA: Kendall/Hunt; 2019

**177**

**Chapter 11**

**Abstract**

management

**1. Introduction**

Environment

Review of Aquatic Biodiversity

*Belda Quetina Mosepele and Ketlhatlogile Mosepele*

strategies to respond to future unexpected events.

**Keywords:** ecosystem perturbations, Okavango delta, resilience, panarchy,

Ecosystems face a major challenge of consistently providing ecological benefits to humanity balanced with biodiversity conservation [1]. This is even more daunting in freshwater wetlands ecosystems, which are not only biological hotspots [2, 3] but are also sources of key ecosystem services that sustain livelihoods globally [2, 4–7]. Services provided by wetlands are not synergistic [2]. According to Maltby and Acreman [6], wetlands are invariably degraded when management enhances the provision of some services like food, which often happens at the expense of other services like regulating services. This scenario is accentuated in tropical areas where, according to Junk [8], politicians prioritise development over environmental protection. Generally, these development priorities are undertaken without any meaningful assessment of the resultant environmental degradation of the wetland [4]. Subsequently, this approach results in wetlands being undervalued "in decisions

Dynamics in the Okavango Delta:

Resilience in a Highly Fluctuating

Wetlands are key ecosystems of high biological diversity that provide valuable ecosystem services. These are particularly important in water stressed semi-arid countries, which enhances their vulnerability to degradation. The Okavango Delta, a key wetland in Botswana, is characterised by dynamic inter and intra specific interactions. There are dynamic biotic and abiotic interactions in the system that enhances its resilience. The flood pulse is the main factor mediating bio-physical dynamics in this system. Despite the various perturbations that have been experienced in the system, the Delta has always been able to absorb them and retain its character at the general ecosystem level. These notwithstanding, there have been some changes at the local scale where the Delta has shifted regimes and entered into altered states as a consequence of either channel or lagoon failure. Management of these systems should ensure that their dynamic characteristics are maintained, and this is enshrined within the panarchy concept. Adopting the resilience framework in natural resources management allows for flexibility in devising management

#### **Chapter 11**
